pores present in cephalic sensory canal system; scapular process greatly enlarged (fig. 8B) Cyprinodon (North, Middle and South America) 44B Males with blackened scales on side; pores absent in cephalic sensory canal system;
scapular process not enlarged Megupsilon
(Nuevo Leon, Mexico) 43B Dorsal fin with more than 15 rays; suborbital bar present; scapular process not promi- nent Jordanella (Florida and the Yucatan Peninsula south to Belize) 42B No dorsal spine; no dorsal ocellus; first pha- ryngobranchial present 45A
45A First pharyngobranchial toothplate present with a patch of unicuspid teeth (fig. 50A);
pectoral fin rays 18-20; inferior pharyn- geals not expanded, teeth with slight shoulders; gill rakers on the first arch 9- 10 Floridichthys (Florida and the Yucatan Peninsula south to Honduras) 45B Cartilaginous first pharyngobranchial lacking teeth (fig. 50B); pectoral fin rays 11-13;
inferior pharyngeals expanded, with closely set villiform teeth (fig. 84); gill rak- ers on the first arch 17 Cualac (San Luis Potosi, Mexico)
row and elongate or narrow and short. Ros- tral cartilage present as a large disc, reduced to a minute disc, or absent.
Interarcual cartilage from base of the first epibranchial to cartilage of second pharyn- gobranchial present or rarely absent; when present large and equal or one-half the length of the first epibranchial. Pharyngobranchial toothplates two, three, and four separate or, three and four fused; or all three fused
to form one large toothplate; pharyngo- branchial teeth unicuspid or tricuspid, often molariform.
Branchiostegal rays three to seven.
Scales in lateral series 24-96, vertebrae 24-54. Oviparous, ovoviviparous or vivipa- rous. Developmental period generally of 10 days or longer. Annual, semiannual, or non- annual reproductive modes. Internal or ex- ternal fertilization.
SUBORDER APLOCHEILOIDEI
DIAGNOSIS: Distinguished from other cy- prinodontiforms by the following uniquely derived characters: orbital rim attached at least on the lower half of the orbit, pelvic fin bases set close together with medial pro- cesses reduced, a broad anterior end of the basihyal, a narrow and twisted lacrimal as- sociated with a narrow preorbital distance, and the posterior extension of the vomer dor- sal to the parasphenoid; distinguished by the following derived characters considered to be convergent in other cyprinodontiforms:
mesethmoid cartilaginous, tubular anterior nans, males always larger than females, and a reduced supraorbital sensory pore system.
COMPOSITION: TWO families, Rivulidae Myers and Aplocheilidae Bleeker, as defined below.
DISTRIBUTION: Pantropical and Old World temperate Laurasian, one family (Rivulidae) New World, the other (Aplocheilidae) Old World, with distributions as detailed below.
FAMILY APLOCHEILIDAE BLEEKER Type Genus Aplocheilus McClelland, 1839.
DIAGNOSIS: Distinguished from all other Cyprinodontiformes in having the supra- cleithrum fused to the posttemporal; pre- maxillary ascending processes tapered pos- teriorly; basihyal reduced to a small triangular ossification; and the interarcual cartilage at- taching directly to the lateral surface of the cartilaginous articular surface of the second pharyngobranchial.
COMPOSITION: Six recognized genera:
Aplocheilus McClelland, Epiplatys Gill with
four subgenera, Pachypanchax Myers, Nothobranchius Peters with four subgenera, Adamas Huber and Aphyosemion Myers with 11 subgenera.
DISTRIBUTION: (fig. 23) Old World; Africa south of the Sahara Desert from the lowlands of southern Mauritania in western Africa, south through Zaire; the northern limit the Niger River then east to western Sudan, south of the Ethiopian Highlands to western Somalia, southward to the coastal lowlands of South Africa; Madagascar; the Sey- chelles; Indian subcontinent and Sri Lanka (Ceylon), eastward through the Indo-Malay- sian Archipelago to Java.
GENUS APLOCHEILUS MCCLELLAND Aplocheilus McClelland, 1839, p. 301 (type
species Esox panchax Hamilton-Buchanan, by original designation).
Panchax Cuvier and Valenciennes, 1846, p. 380 (type species Esox panchax Hamilton-Buchan- an, by original designation).
Haplochilus Agassiz, 1846, p. 24 (proposed as an emendation of Aplocheilus McClelland).
ETYMOLOGY: Aplocheilus from the Greek aplos, meaning single or simple and cheilus, meaning lip, referring to the thin upper and lower jaw margins.
COMPOSITION: Five species: panchax (Hamilton-Buchanan), with seven nominal subspecies; werneri Meinken; lineatus (Cu- vier and Valenciennes); blocki (Arnold); and dayi (Steindachner).
DIAGNOSIS: Distinguished from all other cyprinodontiforms by a derived upper jaw in which the premaxillary ascending processes
overlap in the midline, and the lower jaw is greatly attenuated.
DEFINITION: Anal: Hi, 12-iii, 13; Dorsal:
ii, 6; Pelvic: 6; Pectoral: 14; Caudal: 5, 14, 5. Vertebrae: 13+14. Gill rakers on the an- terior arm of the first arch: 12; Branchioste- gal rays: 6. Scales lateral series: 25-31.
First pleural rib on parapophysis of second vertebra; parapophysis not reduced; a pleu- ral rib often on first hemal spine; hypural plates divided, upper plate often divided in two. Epipleural ribs not bifid.
Anal fin not modified into a gonopodium;
anal fin musculature unmodified; first proxi- mal radial present; middle anal radials pres- ent.
Spermatozeugmata not formed; fertiliza- tion external; development nonannual; ovip- arous. Eggs round.
One dorsal ray articulating with each of the first two dorsal radials; dorsal fin on pos- terior third of body.
Autopterotic fossa normal; lateral ethmoid not expanded medially; not reaching para- sphenoid; parasphenoid not expanded ante- riorly; weakly formed supraoccipital and epiotic processes; neural spine on first ver- tebra; first vertebra articulates with skull via basioccipital and exoccipital condyles; su- praoccipital excluded from formation of fo- ramen magnum; parietals present; nasal not expanded medially.
Mesethmoid cartilaginous; medial pro- cesses of pelvic fin base and ischial process reduced; interarcual cartilage large, attaches laterally to second pharyngobranchial which lacks bony flange; basihyal broad anteriorly, triangular ossification posteriorly; tooth patches on second and third hypobranchials;
teeth on fourth ceratobranchials; dorsal and ventral hypohyal present; anterior extension of anterior ceratohyal ventral to hypohyals;
no uncinate process on fourth epibranchial to articulate with that of third; first epibran- chial narrow at its base. Interhyal ossified;
three ossified basibranchials. Vomer with posterior extension dorsal to parasphenoid.
Lacrimal narrow and twisted carrying dis- tinct sensory canal; dermosphenotic and pre- opercular with distinct sensory canal; pec-
toral girdle lowset; first postcleithrum present; posttemporal with unossified lower limb; posttemporal fused to supracleithrum.
Vomer ossified, dentigerous; medial arm of maxilla twisted with no pronounced dorsal process; ventral arms gently curved toward and abutting rostral cartilage; outer arm nar- row.
Premaxillary ascending processes flat and broad, tapered posteriorly and overlapping in the midline; rostral cartilage large and pen- tagonal; outer arm of premaxilla with alveo- lar process, not indented posteriorly. Liga- ment extending from ventral arms of maxillaries to middle of rostral cartilage; eth- momaxillary ligament present; meniscus present between premaxilla and maxilla.
Dentary not expanded medially, not ro- bust; coronoid process on dentary overlap- ping with that of articular; retroarticular not elongate. Autopalatine with straight head, ventral process not elongate, not reaching quadrate; metapterygoid present.
Orbital rim free dorsally; anterior nares tu- bular; supraorbital sensory pores reduced to a series of neuromasts; seven preopercular pores, four mandibular pores; two or three lacrimal pores.
Males larger than females, often with fin extensions; pigment pattern in many cases composed of several dark crossbars on the sides of the body; all species with a spot at the anterior base of the dorsal fin at least in males; often a darkened caudal margin; mid- dle rays of caudal elongate in juveniles and adults; throat bars present.
No fatty predorsal ridge; caudal scaled for one-third its length; swimbladder extending posteriorly to parhypural.
DISTRIBUTION: Indian subcontinent and Sri Lanka (Ceylon) eastward along the Indo- Malaysian Archipelago to Java.
REMARKS: Members of the genus Aplo- cheilus as defined herein have been most re- cently reviewed by Radda (1973). He fol- lowed Scheel (1972) in treating species of the genus Epiplatys as a subgenus, and there- fore, referred to the species of Aplocheilus as forming a subgenus. In the present study Aplocheilus and Epiplatys are not consid-
ered to be synonyms, rather Aplocheilus is considered to be more closely related to the genus Pachypanchax.
Also, for a variety of reasons, Aplocheilus has long been considered to be the most primitive cyprinodontiform genus. Derived characters it shares with other Old World aplocheiloids refute this hypothesis.
MATERIAL EXAMINED: A. panchax: Gulf of Thailand, CAS 37934 (5*/64); India: Ma- dras, SU 41523 (35); Malaya: SU 32785 (19).
A. lineatus: India: Calicut: SU41516 (3*731).
Aquarium material: AMNH 21498 SW (2+/
2). A. blocki: S. India: Cochin: SU 41513 (14).
GENUS PACHYPANCHAX MYERS Pachypanchax Myers, 1933b, p. 1 (type species
Haplochilus playfairii Gunther, by original designation).
ETYMOLOGY: Pachypanchax from the Greek pachy, meaning robust and Panchax, a synonym of a related genus, referring to the robust appearance of this genus.
TYPES: Seychelles: Haplochilus playfairii Gunther, Syntypes, BMNH 1864.11.15:91- 93(2).
COMPOSITION: TWO species: playfairi (Gunther), and homalanotus (Dumeril).
DIAGNOSIS: Distinguished from other cy- prinodontiforms in having the posterior edge of scales of males angled away from the body in life; from other members of the Aplo- cheilus-Pachy panchax-Epiplatys group in having the hypural plates fused into a hy- pural fan in adults.
DEFINITION: Anal: ii, 15-ii, 16; Dorsal: ii, 9-iii, 9; Pelvic: 6; Pectoral: 19; Caudal: 6, 21, 6; Vertebrae: 13+16. Gill rakers on first arch: 10; Branchiostegal rays: 6. Scales lat- eral series: 25-27.
First pleural rib on parapophysis of second vertebra; parapophysis not reduced; no pleu- ral ribs on hemal spines; hypural plates fused into hypural fan in adults, joint line visible in juveniles. Epipleural ribs not bifid.
Anal fin not modified into a gonopodium;
anal fin musculature unmodified; first proxi-
mal radial present; middle anal radials pres- ent.
Spermatozeugmata not formed; fertiliza- tion external; development nonannual; ovip- arous. Eggs round.
One dorsal ray articulating with each of the first two dorsal radials; dorsal fin on pos- terior third of body.
Autopterotic fossa normal; lateral ethmoid not expanded medially, not reaching para- sphenoid; parasphenoid not expanded ante- riorly; weakly formed supraoccipital and epiotic processes; neural spine on first ver- tebra; first vertebra articulates with skull via basioccipital and exoccipital condyles; su- praoccipital excluded from formation of fo- ramen magnum; parietals present; nasals not expanded medially.
Mesethmoid cartilaginous; medial pro- cesses of pelvic fin base and ischial process reduced; interarcual cartilage large, attaches laterally to second pharyngobranchial which lacks bony flange; basihyal broad anteriorly, triangular ossification posteriorly; tooth patches on second and third hypobranchials;
teeth on fourth ceratobranchials; dorsal and ventral hypohyal present; anterior extension of anterior ceratohyal ventral to hypohyals;
no uncinate process on fourth epibranchial to articulate with that of third; first epibranch- ial narrow at its base. Interhyal ossified;
three ossified basibranchials. Vomer with posterior extension dorsal to parasphenoid.
Lacrimal narrow and twisted carrying dis- tinct sensory canal; dermosphenotic and pre- opercular with distinct sensory canal; pec- toral girdle lowset; first postcleithrum present; posttemporal with unossified lower limb; posttemporal fused to supracleithrum.
Vomer ossified, dentigerous; medial arm of maxilla twisted with no pronounced dorsal process; ventral arms gently curved toward and abutting rostral cartilage; outer arm nar- row.
Premaxillary ascending processes flat and broad, tapered posteriorly, not overlapping in the midline; rostral cartilage large and pen- tagonal; outer arm of premaxilla with alveo- lar process, not indented posteriorly. Liga- ment extending from ventral arms of
maxillaries to middle of rostral cartilage; eth- momaxillary ligament present; meniscus present between premaxilla and maxilla.
Dentary not expanded medially, not ro- bust; coronoid process on dentary overlap- ping with that of articular; retroarticular not elongate. Autopalatine with straight head, ventral process not elongate, not reaching quadrate; metapterygoid present.
Orbital rim free dorsally; anterior naris tu- bular; supraorbital sensory pores reduced to a series of neuromasts; seven preopercular pores, four mandibular pores; two or three lacrimal pores.
Males larger than females, never with fin extensions; pigment pattern not consisting of crossbars; males and females with faint red reticulations; juveniles and females with spot at anterior base of the dorsal fin (in play- fairi); a darkened caudal and anal margin in males of play>j"aid; middle rays of caudal nev- er elongate; throat bars present.
No fatty predorsal ridge; caudal scaled for at least one-third its length; swimbladder ex- tending posteriorly to parhypural.
DISTRIBUTION: Madagascar, the Sey- chelles, coastal lowlands of eastern Mozam- bique, and Zanzibar north of the Zambezi River.
REMARKS: The genus Pachypanchax is considered to contain two species: playfairi, the type, and homalanotus. However, a specimen of homalanotus was not available for osteological examination, therefore, it is included with caution in the genus.
MATERIAL EXAMINED: P. playfairi: Sey- chelles: Syntypes as listed above, AMNH 20637 (4*/17); Zanzibar: AMNH 20701 (4*/
19); Aquarium material: AMNH 38413 (3*/
3); P. homalanotus: Aquarium material: SU 52679(1).
GENUS EPIPLATYS GILL
Epiplatys Gill, 1862, p. 136 (type species Haplo- chilus sexfasciatus Gill, by original designa- tion).
ETYMOLOGY: Epiplatys from the Greek epi, meaning above, and platys meaning flat,
referring to the flattened dorsal aspect of the skull.
TYPES: Gaboon: Haplochilus sexfasciatus Gill, Types, ANSP 7129 to 7141.
COMPOSITION: Over 50 nominal species and subspecies as listed in Lazara (1979).
DIAGNOSIS: Distinguished from other members of the Aplocheilus-Pachypanchax- Epiplatys group by the following primitive characters found derived in Aplocheilus and Pachypanchax: posttemporal with an ossi- fied lower limb; orbital rim completely at- tached; no teeth on the hypobranchials; no darkened caudal margin, and no dorsal ocel- lus.
DEFINITION: Anal: iv, 12; Dorsal: iv, 7;
Pelvic: 6; Pectoral: 17; Caudal: 6, 20, 6; Ver- tebrae: 11 + 14. Gill rakers on the anterior arm of the first arch: 8; Branchiostegal rays:
6. Scales lateral series: 26-30.
First pleural rib on parapophysis of second vertebra; parapophysis not reduced; a pleu- ral rib often on first hemal spine; hypural plates divided, upper plate often divided in two. Epipleural ribs not bifid.
Anal fin not modified into a gonopodium;
anal fin musculature unmodified; first proxi- mal radial present; middle anal radials pres- ent.
Spermatozeugmata not formed; fertiliza- tion external; development nonannual; ovip- arous. Eggs round.
One dorsal ray articulating with each of the first two dorsal radials; dorsal fin on pos- terior third of body.
Autopterotic fossa normal; lateral ethmoid not expanded medially, not reaching para- sphenoid; parasphenoid not expanded ante- riorly; weakly formed supraoccipital and epiotic processes; neural spine on first ver- tebra; first vertebra articulates with skull via basioccipital and exoccipital condyles; su- praoccipital excluded from formation of fo- ramen magnum; parietals present; nasals not expanded medially.
Mesethmoid cartilaginous; medial pro- cesses of pelvic fin base and ischial process reduced; interarcual cartilage large, attaches laterally to second pharyngobranchial which lacks bony flange; basihyal broad anteriorly,
triangular ossification posteriorly; no tooth patches on second and third hypobranchials;
teeth on fourth ceratobranchials; dorsal and ventral hypohyal present; anterior extension of anterior ceratohyal ventral to hypohyals;
no uncinate process on fourth epibranchial to articulate with that of third; first epibran- chial narrow at its base. Interhyal ossified;
three ossified basibranchials. Vomer with posterior extension dorsal to parasphenoid.
Lacrimal narrow and twisted carrying dis- tinct sensory canal; dermosphenotic and pre- opercular with distinct sensory canal; pec- toral girdle lowset; first postcleithrum present; posttemporal with ossified lower limb; posttemporal fused to supracleithrum.
Vomer ossified, dentigerous; medial arm of maxilla twisted with no pronounced dorsal process; ventral arms gently curved toward and abutting rostral cartilage; outer arm nar- row.
Premaxillary ascending processes flat and broad, tapered posteriorly, not overlapping in the midline; rostral cartilage large and pen- tagonal; outer arm of premaxilla with alveo- lar process, not indented posteriorly. Liga- ment extending from ventral arms of maxillaries to middle of rostral cartilage; eth- momaxillary ligament present; meniscus present between premaxilla and maxilla.
Dentary not expanded medially, not ro- bust; coronoid process on dentary not over- lapping with that of articular; retroarticular not elongate. Autopalatine with straight head, ventral process not elongate, not reaching quadrate; metapterygoid present.
Orbital rim attached; anterior naris tubu- lar; supraorbital sensory pores reduced to a series of neuromasts; seven preopercular pores, three mandibular pores; two or three lacrimal pores.
Males larger than females, often with fin extensions; pigment pattern often composed of several dark crossbars on the sides of the body; no species with a spot at the anterior base of the dorsal fin; middle rays of caudal elongate in juveniles and adults; throat bars present.
No fatty predorsal ridge; caudal scaled for at least one-third its length; swimbladder ex-
tending posteriorly to first or second hemal spine.
DISTRIBUTION: West and central Africa:
eastward from Senegal, northern limit the Niger River, eastward to the Ethiopian High- lands, southward west of the Rift lakes to the Katanga Plateau, then eastward to the coast.
REMARKS: The genus Epiplatys as consti- tuted in this study is paraphyletic. However, rather than create new generic names for subgroups of the genus which could not be incorporated into an overall phylogenetic scheme, the species group names which al- ready exist are treated as subgenera of the genus Epiplatys until such time that they may be defined and their relationships deter- mined. These subgenera are the following:
SUBGENUS LYCOCYPRINUS PETERS Lycocyprinus Peters, 1868, p. 146 (type species
Epiplatys dageti Poll by monotypy).
SUBGENUS PAREPIPLATYS CLAUSEN Parepiplatys Clausen, 1967, p. 28 (type species
Haplochilus grahami Boulenger, by original designation [proposed as a subgenusj).
SUBGENUS PSEUDEPIPLATYS CLAUSEN Pseudepiplatys Clausen, 1967, p. 30 (type species
Haplochilus annulatus Boulenger, by original designation [proposed as a subgenus]).
SUBGENUS APHYOPLATYS CLAUSEN Aphyoplatys Clausen, 1967, p. 32 (type species
Epiplatys duboisi Poll, by original designation [proposed as a subgenus]).
MATERIAL EXAMINED: E. sexfasciatus:
Dahomey, Iquidi R: USNM 218752 (2*721);
E. chaperi: Ghana: SU 64709 (1*712); E.fas- ciolatus: Liberia: Tchien: AMNH 32735 (5*7 10); E. bifasciatus: Nigeria (Aquarium ma- terial) AMNH 21866SW (1+/1). E. senegal- ensis: Nigeria (Aquarium material) AMNH 21867 (1+/1); E. dageti: Ghana: SU 64640 (29). Haplochilus annulatus: Type: Sierra Leone: BMNH 1914. 12. 9: 5-6.
GENUS APHYOSEMION MYERS
Aphyosemion Myers, 1924b, p. 2 (type species Aphysemion castaneum Myers, by original designation).
ETYMOLOGY: Aphyosemion from the Greek aphyos, meaning small and semion meaning flag or banner referring to the lyre- shaped caudal fin of males.
TYPES: Zaire (Congo): Aphyosemion cas- taneum Myers, Type AMNH 8337 (1).
COMPOSITION: Approximately 60 species as listed in Lazara (1979).
DIAGNOSIS: Distinguished from other members of the Aphyosemion-Nothobran- chius group by being nonannual, having a dorsal fin of seven to 14 rays which is situ- ated no farther anteriorly than opposite the first quarter of the anal fin orgin, and pos- sessing a swimbladder extending posteriorly to the first one or two hemal spines.
DEFINITION: Anal: iii, 12; Dorsal: i, 6-ii, 12; Pelvic: 6; Pectoral: 16-17; Caudal: 8, 13, 8; Vertebrae: 13 + 15. Gill rakers on anterior arm of the first arch: 8, 9; Branchiostegal rays: 6. Scales Lateral Series: 29-33.
First pleural rib on parapophysis of second vertebra; parapophysis not reduced; no pleu- ral rib on first hemal spine; hypural plates divided, upper plate never divided in two.
Epipleural ribs bifid.
Anal fin not modified into a gonopodium;
anal fin musculature unmodified; first proxi- mal radial present; middle anal radials pres- ent.
Spermatozeugmata not formed; fertiliza- tion external; development nonannual; ovip- arous. Eggs round.
One dorsal ray articulating with each of the first two dorsal radials; dorsal fin on pos- terior third of body.
Autopterotic fossa normal; lateral ethmoid not expanded medially, not reaching para- sphenoid; parasphenoid not expanded ante- riorly; weakly formed supraoccipital and epiotic processes; neural spine on first ver- tebra; first vertebra articulates with skull via basioccipital and exoccipital condyles; su- praoccipital excluded from formation of fo- ramen magnum; parietals present; nasals not expanded medially.
Mesethmoid cartilaginous; medial pro- cesses of pelvic fin base and ischial process reduced; interarcual cartilage large, attaches laterally to second pharyngobranchial which lacks bony flange; basihyal broad anteriorly, triangular ossification posteriorly; no tooth patches on second and third hypobranchials;
teeth on fourth ceratobranchials; dorsal and ventral hypohyal present; anterior extension of anterior ceratohyal ventral to hypohyals;
uncinate process on fourth epibranchial ar- ticulates with that of third; first epibranchial narrow at its base. Interhyal ossified; three ossified basibranchials. Vomer with atten- uate posterior extension dorsal to parasphe- noid.
Lacrimal narrow and twisted carrying dis- tinct sensory canal; dermosphenotic and pre- opercular with distinct sensory canal; pec- toral girdle lowset; first postcleithrum present; posttemporal with ossified lower limb; posttemporal fused to supracleithrum.
Vomer ossified, dentigerous; medial arm of maxilla twisted with no pronounced dorsal process; ventral arms gently curved toward and abutting rostral cartilage; outer arm nar- row.
Premaxillary ascending processes flat and broad, tapered posteriorly not overlapping in the midline; rostral cartilage large and pen- tagonal; outer arm of premaxilla with alveo- lar process, not indented posteriorly. Liga- ment extending from ventral arms of maxillaries to middle of rostral cartilage; eth- momaxillary ligament present; meniscus present between premaxilla and maxilla.
Dentary not expanded medially, not ro- bust; coronoid process on dentary not over- lapping with that of articular; retroarticular not elongate. Autopalatine with straight head, ventral process not elongate, not reaching quadrate; metapterygoid present.
Orbital rim attached; anterior nans tubu- lar; supraorbital sensory pores reduced to a series of neuromasts; seven preopercular pores, four mandibular pores; two or three lacrimal pores.
Males larger than females, often with fin extensions; pigment pattern rarely composed of several dark crossbars on the sides of the body; no species with a spot at the anterior